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The Relationships of the Euparkeriidae and the Rise of Archosauria

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The Relationships of the Euparkeriidae and the Rise of Archosauria Downloaded from http://rsos.royalsocietypublishing.org/ on March 29, 2017 The relationships of the Euparkeriidae and the rise rsos.royalsocietypublishing.org of Archosauria 1,2 Research Roland B. Sookias 1School of Geography, Earth and Environmental Sciences, University of Birmingham, Cite this article: Sookias RB. 2016 The Edgbaston, Birmingham B15 2TT, UK relationships of the Euparkeriidae and the rise 2GeoBio-Center, Ludwig-Maximilians-Universität München, Richard-Wagner-Straße of Archosauria. R. Soc. open sci. 3: 150674. 10, 80333 Munich, Germany http://dx.doi.org/10.1098/rsos.150674 For the first time, a phylogenetic analysis including all putative euparkeriid taxa is conducted, using a large data matrix Received: 8 December 2015 analysed with maximum parsimony and Bayesian analysis. Accepted: 19 February 2016 Using parsimony, the putative euparkeriid Dorosuchus neoetus from Russia is the sister taxon to Archosauria + Phytosauria. Euparkeria capensis is placed one node further from the crown, and forms a euparkeriid clade with the Chinese taxa Halazhaisuchus qiaoensis and ‘Turfanosuchus shageduensis’and Subject Category: the Polish taxon Osmolskina czatkowicensis. Using Bayesian Biology (whole organism) methods, Osmolskina and Halazhaisuchus are sister taxa within Euparkeriidae, in turn sister to ‘Turfanosuchus shageduensis’and Subject Areas: then Euparkeria capensis. Dorosuchus is placed in a polytomy + evolution/palaeontology with Euparkeriidae and Archosauria Phytosauria. Although conclusions remain tentative owing to low node support and incompleteness, a broad phylogenetic position close Keywords: to the base of Archosauria is confirmed for all putative Euparkeriidae, Archosauria, Triassic, euparkeriids, and the ancestor of Archosauria +Phytosauria Archosauriformes, phylogeny is optimized as similar to euparkeriids in its morphology. Ecomorphological characters and traits are optimized onto the maximum parsimony strict consensus phylogeny presented using squared change parsimony. This optimization indicates Author for correspondence: that the ancestral archosaur was probably similar in many Roland B. Sookias respects to euparkeriids, being relatively small, terrestrial, e-mail: [email protected] carnivorous and showing relatively cursorial limb morphology; this Bauplan may have underlain the exceptional radiaton and success of crown Archosauria. 1. Introduction Archosauria, the diapsid clade represented today by birds and crocodilians and including the extinct dinosaurs, is highly speciose (with over 9000 species of extant bird and crocodilian [1]). Archosaurs filled most major terrestrial ecological niches for Electronic supplementary material is available over 150 million years [2–7], from the Middle Triassic to the end at http://dx.doi.org/10.1098/rsos.150674 or via of the Cretaceous. The ‘rise’ of the archosaurs to this position of http://rsos.royalsocietypublishing.org. 2016 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited. Downloaded from http://rsos.royalsocietypublishing.org/ on March 29, 2017 ecological dominance took place coincident with the extinction or decline of many therapsid taxa at 2 the end of the Permian and during the course of the Early–Middle Triassic, which had previously rsos.royalsocietypublishing.org filled most large-bodied terrestrial niches [2,3,8–14]. The rise of the archosaurs is one of the landmark ................................................ terrestrial faunal transitions and is an outstanding example of a large-scale adaptive radiation in the fossil record [2,15,16], with archosaurs diversifying into carnivorous, herbivorous, aquatic, terrestrial and volant forms of greatly varying sizes [2,14–17]. Understanding this adaptive radiation requires a thorough knowledge of archosaur phylogeny, of the morphological changes seen during this radiation and the sequence of these changes. The archosaur radiation must also be seen in its wider context as part of a radiation of archosauromorphs (those taxa more closely related to crown Archosauria than crown Lepidosauria), with aspects of the archosaur body plan beginning to develop within this stem lineage, and setting the stage for the unprecedented success of crown Archosauria. One such example is the development of cursorial locomotion [18], which may have R. Soc. open sci. allowed archosaurs to radiate into carnivorous niches following the extinction of therapsid carnivores. The family Euparkeriidae has been historically composed of a number of small, gracile archosauriform taxa that have often been placed immediately outside or close to the base of Archosauria [19](figure 1). They have often been discussed as potentially representing a morphology 3 very close to that of the ancestral archosaur [24], although this idea has not been quantitatively : 150674 investigated. However, irrespective of whether it represented a true phylogenetically intermediate step, the gracile, cursorial morphology of euparkeriids is roughly intermediate between more ‘sprawling’ early archosauromorph taxa and fully erect, and often bipedal, archosaurs [14,18,25]. Furthermore, given their phylogenetic position alone, these animals have the potential to shed light on the patterns seen during the archosaur radiation, and the factors that underlay archosaur success. The only taxon to be assigned with certainty to Euparkeriidae however is Euparkeria capensis, and the monophyly of the family has remained largely untested until recently. Here, for the first time, a phylogenetic analysis is conducted including all putative euparkeriids and a representative subset of stem and crown archosaurs, incorporating both new characters and those taken from previous analyses. 2. Previous phylogenetic work 2.1. Composition of Euparkeriidae Following recent revisions, there are four valid species that may represent euparkeriids: Euparkeria capensis [26,27] (the type genus and species of the family Euparkeriidae), Halazhaisuchus qiaoensis (see [28]) and Osmolskina czatkowicensis (see [29,30]). Moreover, the holotype specimen of the nomen dubium ‘Turfanosuchus shageduensis’ has also been considered possibly referable to the clade [28]. Recent phylogenetic work has recovered Halazhaisuchus qiaoensis,‘Turfanosuchus shageduensis’andEuparkeria capensis within a euparkeriid clade, with the former two taxa being sister taxa to the exclusion of Euparkeria capensis [28], whereas Dorosuchus neoetus has been placed outside Euparkeriidae, one node closer to the crown, as the sister taxon to Archosauria + Phytosauria [31]. The only previous analysis to include Osmolskina czatkowicensis recovered it further down the archosaur stem than Euparkeria capensis [22], although this analysis was carried out before full description of the former taxon. No previous phylogenetic analysis has simultaneously included all of these species. 2.2. Position of Euparkeriidae Most recent work has placed Euparkeria capensis and other euparkeriids close to the base of, but outside, Archosauria (see for example the summaries in [32,33], the placement hypothesized by Borsuk- Białynicka & Evans [30], and the phylogenetic analyses of Benton & Clark [34], Sereno & Arcucci [35], Sereno [36], Juul [37], Bennett [38], Benton [39], Parker & Barton [40], Nesbitt et al.[41], Dilkes & Sues [21], Nesbitt [23], Brusatte et al.[2], Ezcurra et al.[22], Desojo et al.[42], Dilkes & Arcucci [43], Schoch & Sues [44], Sookias et al.[28,31], Parrish [45]; figure 1b–d). This contrasts with the placement of Euparkeria capensis within the crown, as the sister taxon to Ornithosuchidae + Ornithodira (within ‘Ornithosuchia’), found in an early analysis by Gauthier [20](figure 1a). Less consensus has been reached regarding the relationships between euparkeriids and other stem archosaurs. Several analyses have placed Euparkeria capensis as the sister taxon to Archosauria [34,38,41,45] or Phytosauria + Archosauria [23,28,31]. However, many previous analyses have also placed proterochampsids, or proterochampsids + doswelliids, closer to the crown than is Euparkeria capensis Downloaded from http://rsos.royalsocietypublishing.org/ on March 29, 2017 (a)(b) PROTEROSUCHIDAE Mesosuchus browni 3 rsos.royalsocietypublishing.org ................................................ Prolacerta broomi ERYTHROSUCHIDAE Proterosuchus fergusi Euparkeria capensis PROTEROCHAMPSIDAE Erythrosuchus africanus PSEUDOSUCHIA Doswellia kaltenbachi R. Soc. open sci. PROTEROCHAMPSIDAE ARCHOSAURIA Euparkeria capensis Turfanosuchus dabanensis 3 Yonghesuchus sangbiensis : 150674 ORNITHOSUCHIA ORNITHOSUCHIDAE PSEUDOSUCHIA ORNITHODIRA ARCHOSAURIA ORNITHODIRA (c)(d) Mesosuchus browni Mesosuchus browni Prolacerta broomi Prolacerta broomi Proterosuchus fergusi Fugusuchus hejiapanensis Proterosuchus fergusi Sarmatosuchus otschevi Erythrosuchus africanus Osmolskina czatkowicensis Vancleavea campi Koilamasuchus gonzalezdiazi ERYTHROSUCHIDAE PROTEROCHAMPSIA Euparkeria capensis Euparkeria capensis Chanaresuchus bonapartei Vancleavea campi PHYTOSAURIA Doswellia kaltenbachi PSEUDOSUCHIA PSEUDOSUCHIA ARCHOSAURIA ARCHOSAURIA AVEMETATARSALIA AVEMETATARSALIA Figure 1. Previous phylogenetic positions found for Euparkeria capensis.(a)[20]; (b)[21]; (c)[22]and(d)[23]. [2,22,35–37,42] (Benton [39] presents this relationship, but the opposite topology is no less parsimonious; figure 1b,c). All previous phylogenetic work
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